Английская Википедия:Irish elk

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Шаблон:Short description Шаблон:Use Hiberno-English Шаблон:Use dmy dates Шаблон:Speciesbox

The Irish elk (Megaloceros giganteus),[1][2] also called the giant deer or Irish deer, is an extinct species of deer in the genus Megaloceros and is one of the largest deer that ever lived. Its range extended across Eurasia during the Pleistocene, from Ireland (where it is known from abundant remains found in bogs) to Lake Baikal in Siberia. The most recent remains of the species have been radiocarbon dated to about 7,700 years ago in western Russia.[3][4] Its antlers, which can span Шаблон:Convert across are the largest known of any deer.[5] It is not closely related to either living species called the elk, with it being widely agreed that its closest living relatives are fallow deer (Dama).[5][6][7][8]

Taxonomy

Research history

Файл:Megaloceros 1856.png
Skeletal reconstruction from 1856

The first scientific descriptions of the animal's remains were made by Irish physician Thomas Molyneux in 1695, who identified large antlers from Dardistown—which were apparently commonly unearthed in Ireland—as belonging to the elk (known as the moose in North America), concluding that it was once abundant on the island.[9] It was first formally named as Alce gigantea by Johann Friedrich Blumenbach in his Handbuch der Naturgeschichte in 1799,[10] with Alce being a variant of Alces, the Latin name for the elk. The original Blumenbach's description of Alce gigantea provides rather scant information about the species, specifying only that this particular kind of "fossil elk" comes from Ireland and is characterized by immense body size. According to Blumenbach,[10] the distance between summits of giant deer antlers may attain 14 feet (approximately 4.4 m). This particular feature mentioned by Blumenbach permitted to Roman Croitor to identify the type specimen of giant deer [11] that was figured and described for the first time in Louthiana of Thomas Wright.[12] The holotype of Megaloceros giganteus (Blumenbach, 1799) is a well-preserved male skull with exceptionally large antlers found in Dunleer environs (County Louth, Ireland).[11] The type specimen of giant deer is currently exposed in Barmeath Castle where Thomas Wright first saw and described it.[11]

French scientist Georges Cuvier documented in 1812 that the Irish elk did not belong to any species of mammal currently living, declaring it "le plus célèbre de tous les ruminans fossiles" (the most famous of all fossil ruminants).[13] In 1827 Joshua Brookes, in a listing of his zoological collection, named the new genus Megaloceros (spelled Megalocerus in the earlier editions) in the following passage:[14][15]

Шаблон:Blockquote

The etymology being from Greek: Шаблон:Lang Шаблон:Transliteration "great" + Шаблон:Lang Шаблон:Transliteration "horn, antler".[16] The type and only species named in the description being Megaloceros antiquorum, based on Irish remains now considered to belong to M. giganteus, making the former a junior synonym. The original description was considered by Adrian Lister in 1987 to be inadequate for a taxonomic definition.[2] In 1828 Brookes published an expanded list in the form of a catalogue for an upcoming auction, which included the Latin phrase "Cornibus deciduis palmatis" as a description of the remains. The 1828 publication was approved by International Commission on Zoological Nomenclature (ICZN) in 1977 as an available publication for the basis of zoological nomenclature.[2] Adrian Lister in 1987 judged that "the phase "Cornibus deciduis palmatis" constitutes a definition sufficient under the [International Code of Zoological Nomenclature] (article 12) to validate Megalocerus."[2] The original spelling of Megalocerus was never used after its original publication.[15]

Файл:Extinct monsters BHL20699843.jpg
Outdated 1897 reconstruction of doe and stag Irish elk by Joseph Smit

In 1844 Richard Owen named another synonym of the Irish elk, including it within the newly named subgenus Megaceros, Cervus (Megaceros) hibernicus. This has been suggested to be derived from another junior synonym of the Irish elk described by J. Hart in 1825, Cervus megaceros.[2] Despite being a junior synonym, Megaloceros remained in obscurity and Megaceros became the common genus name for the taxon.[15] The combination "Megaceros giganteus" was in use by 1871.[17] George Gaylord Simpson in 1945 revived the original Megaloceros name, which became progressively more widely used, until a taxonomic decision in 1989 by the ICZN confirmed the priority of Megaloceros over Megaceros, and Megaloceros to be the correct spelling.[15][18]

Before the 20th century, the Irish elk, having evolved from smaller ancestors with smaller antlers, was taken as a prime example of orthogenesis (directed evolution), an evolutionary mechanism opposed to Darwinian evolution in which the successive species within the lineage become increasingly modified in a single undeviating direction, evolution proceeding in a straight line void of natural selection. Orthogenesis was claimed to have caused an evolutionary trajectory towards antlers that became larger and larger, eventually causing the species' extinction because the antlers grew to sizes which inhibited proper feeding habits and caused the animal to become trapped in tree branches.[6] In the 1930s, orthogenesis was disputed by Darwinians led by Julian Huxley, who noted that antler size was not grossly large, and was proportional to body size.[19][20] The currently favoured view is that sexual selection was the driving force behind the large antlers rather than orthogenesis or natural selection.[20]

Evolution

Файл:Megaceros giganteus antecedens.JPG
Skull of M. g. antecedens

M. giganteus belongs to the genus Megaloceros. Megaloceros has often been placed into the tribe Megacerini, alongside other genera often collectively referred to as "giant deer", like Sinomegaceros and Praemegaceros.[21] The taxonomy of giant deer lacks consensus, with genus names used for species varying substantially between authors.[22][23] The earliest possible record of the genus is a partial antler from the Early Pleistocene MN 17 (2.5–1.8 Ma) of Stavropol Krai in the North Caucasus of Russia, which were given the name of M. stavropolensis in 2016,[24] however this species has been subsequently suggested to belong to Arvernoceros.[22][23] or Sinomegaceros.[11] The oldest generally accepted records of the genus are from the late Early Pleistocene.[25] Other species often considered to belong to Megaloceros include the reindeer sized M. savini, which is known from early Middle Pleistocene (~700,000–450,000 years ago) localities in England, France, Spain and Germany, and the more recently described species M. novocarthaginiensis, which is known from late Early Pleistocene (0.9–0.8 Ma) localities in Spain, and the small M. matritensis endemic to the Iberian peninsula during the late Middle Pleistocene (~400,000 to 250,000 years ago), which overlaps chronologically with the earliest M. giganteus records. Jan van der Made proposed M. novocarthaginiensis , M. savini and M. matritensis to be sequential chronospecies, due to shared morphological characteristics not found in M. giganteus and gradual transition of morphological characters through time.[22] M. savini and related species have also been suggested to comprise the separate genus Praedama by other authors. Roman Croitor has suggested closer affinities to Eucladoceros for M. savini and related species.[23]

Файл:Knight Megaloceros.jpg
Outdated 1906 restoration by Charles R. Knight

The origin of M. giganteus remains unclear, and appears to lie outside Western Europe.[22] Jan van der Made has suggested that remains of an indeterminate Megaloceros species from the late Early Pleistocene (~1.2 Ma) of Libakos in Greece are closer to M. giganteus than the M. novocarthaginiensis-savini-matritensis lineage due to the shared molarisation of the lower fourth premolar (P4).[22] Croitor has suggested that M. giganteus is closely related to what was originally described as Dama clactoniana mugharensis (which he proposes be named Megaloceros mugharensis) from the Middle Pleistocene of Tabun Cave in Israel, due to similarities in the antlers, molars and premolars.[23] The earliest possible records of M. giganteus comes from Homersfield, England thought to be about 450,000 years ago—though the dating is uncertain.[26] The oldest securely dated Middle Pleistocene records are those from Hoxne, England, which have been dated to Marine Isotope Stage 11 (424,000 to 374,000 years ago),[27][22] other Middle Pleistocene early records include Steinheim an der Murr, Germany, (classified as M. g. antecedens) about 400,000–300,000 years ago and Swanscombe, England.[26][22] Most remains of the Irish elk are known from the Late Pleistocene. A large proportion of the known remains of M. giganteus are from Ireland, which mostly date to the Allerød oscillation near the end of the Late Pleistocene around 13,000 years ago. Over 100 individuals have been found in Ballybetagh Bog near Dublin.[28]

It has been historically thought that, because both have palmated antlers, the Irish elk and fallow deer (Dama spp.) are closely related, this is supported by several other morphological similarities, including the lack of upper canines, proportionally long braincase and nasal bones, and proportionally short front portion of the skull.[23] In 2005, two fragments of mitochondrial DNA (mtDNA) from the cytochrome b gene were extracted and sequenced from 4 antlers and a bone, the mtDNA found that the Irish elk was nested within Cervus, and were inside the clade containing living red deer (Cervus elaphus). Based on this, the authors suggested that the Irish elk and red deer interbred.[29] However, another study from the same year in the journal Nature utilising both fragmentary mitochondrial DNA and morphological data found that the Irish elk was indeed most closely related to Dama.[5] The close relationship with Dama was supported by another cytochrome b study in 2006,[6] a 2015 study involving the full mitochondrial genome,[7] and by a 2017 morphological analysis of the bony labyrinth.[8] The 2006 and 2017 studies also directly suggest that the results of the 2005 cytochrome b paper were the result of DNA contamination.[6][8]

Cladogram of Cervidae based on mitochondrial DNA:[30]

Шаблон:Clade

A study of mitochondrial genomes from Sinomegaceros from the Late Pleistocene of East Asia found that the mitochondrial genomes of Megaloceros giganteus were nested within those of Sinomegaceros, suggesting that the two lineages interbred after their initial split. Cladogram of Megaloceros and Sinomegaceros mitochondrial genomes following Xiao et al. 2023.[31]

Шаблон:Clade

Description

Файл:Megaloceros.jpg
Life restoration

The Irish elk stood about Шаблон:Cvt tall at the shoulders,[5] and had large palmate (flat and broad) antlers,[32] the largest known of any known deer, with the largest specimens reaching over Шаблон:Cvt from tip to tip[5] (though it is rare for specimens to exceed Шаблон:Convert across[11]) and Шаблон:Cvt in weight.[33] The antlers are considerably larger than living moose, being on average over twice the volume of moose antlers.[30] For body size, at about Шаблон:Cvt and up to Шаблон:Cvt or more,[34][33][35] the Irish elk was the heaviest known cervine ("Old World deer");[5] and tied with the extant Alaska moose (Alces alces gigas) as the third largest known deer, after the extinct Cervalces latifrons and Cervalces scotti.[34][33] Nonetheless, compared to Alces, Irish elk appear to have had a more robust skeleton, with older and more mature Alces skeletons bearing some resemblance to those of prime Irish elk, and younger Irish elk resembling prime Alces. Likely due to different social structures, the Irish elk exhibits more marked sexual dimorphism than Alces, with Irish elk bucks being notably larger than does.[36] In total, Irish elk bucks may have ranged from Шаблон:Cvt, with an average of Шаблон:Cvt, and does may have been relatively large, about 80% of buck size, or Шаблон:Cvt on average.[37] The distinguishing characters of M. giganteus include concave frontals, proportionally long braincase, proportionally short front section of the skull (orbitofrontal region), alongside the absence of upper canines and the molarisation of the lower fourth premolar (P4). The skull and mandible of the Irish elk exhibit substantial thickening (pachyostosis), with the early and complete obliteration of cranial sutures.[23]

Based on Upper Palaeolithic cave paintings, the Irish elk seems to have had overall light colouration, with a dark stripe running along the back, a stripe on either side from shoulder to haunch, a dark collar on the throat and a chinstrap, and a dark hump on the withers (between the shoulder blades). In 1989, American palaeontologist Dale Guthrie suggested that, like bison, the hump allowed a higher hinging action of the front legs to increase stride length while running. Valerius Geist suggested that the hump may have also been used to store fat. Localising fat rather than evenly distributing it may have prevented overheating while running or in rut during the summer.[37]

Habitat

It was not exclusive to Ireland. Rather, it was so named because the most well-known and best-preserved fossil specimens have been found in lake sediments and peat bogs in Ireland. The Irish elk had a far-reaching range, extending from the Atlantic Ocean in the West to Lake Baikal in the East. They do not appear to have extended northward onto the open mammoth steppe, rather keeping to the boreal steppe-woodland environments, which consisted of scattered spruce and pine, as well as low-lying herbs and shrubs including grasses, sedges, Ephedra, Artemisia and Chenopodiaceae.[4]

Palaeobiology

Physiology

Файл:Megaloceros cougnac.svg
Drawing of cave art from Grotte de Cougnac, France showing coloured shoulder hump and lines. c. 25,000 to 19,000 years old[38]

In 1998, Canadian biologist Valerius Geist hypothesised that the Irish elk was cursorial (adapted for running and stamina). He noted that the Irish elk physically resembled reindeer. The body proportions of the Irish elk are similar to those of the cursorial addax, oryx, and saiga antelope. These include the relatively short legs, the long front legs nearly as long as the hind legs, and a robust cylindrical body. Cursorial saiga, gnus, and reindeer have a top speed of over Шаблон:Cvt, and can maintain high speeds for up to 15 minutes.[37]

Reproduction

Шаблон:Multiple image

At Ballybetagh Bog, over 100 Irish elk individuals were found, all small antlered bucks. This indicates that bucks and does segregated during at least winter and spring. Many modern deer species do this partly because males and females have different nutritional requirements and need to consume different types of plants. Segregation would also imply a polygynous society, with stags fighting for control over harems during rut. Because most of the individuals found were juvenile or geriatric and were likely suffering from malnutrition, they probably died from winterkill. Most Irish elk specimens known may have died from winterkill, and winterkill is the highest source of mortality among many modern deer species. Bucks generally suffer higher mortality rates because they eat little during the autumn rut.[39] For rut, a lean stag normally Шаблон:Cvt may have fattened up to Шаблон:Cvt, and would burn through the extra fat over the next month.[37]

Assuming a similar response to starvation as red deer, a large, healthy Irish elk stag with Шаблон:Cvt antlers would have had Шаблон:Cvt antlers under poor conditions;[13][33] and an average sized Irish elk stag with Шаблон:Cvt antlers would have had Шаблон:Cvt antlers under poorer conditions,[40] similar sizes to the moose. A similar change in a typical Irish elk population with prime stags having Шаблон:Cvt antlers would result in antler weights of Шаблон:Cvt or less in worsening climatic conditions. This is within the range of present-day wapiti/red deer (Cervus spp.) antler weights.[37] Irish elk antlers vary widely in form depending upon the habitat, such as a compact, upright shape in closed forest environments.[40] Irish elk likely shed their antlers and re-grew a new pair during mating season. Antlers generally require high amounts of calcium and phosphate, especially those for stags which have larger structures, and the massive antlers of Irish elk may have required much greater quantities. Stags typically meet these requirements in part from their bones, suffering from a condition similar to osteoporosis while the antlers are growing, and replenishing them from food plants after the antlers have grown in or reclaiming nutrients from shed antlers.[33]

The large antlers have generally been explained as being used for male-male battle during mating season.[41] They may have also been used for display,[13] to attract females and assert dominance against rival males.[39] A finite element analysis of the antlers suggested that during fighting, the antlers were likely to interlock around the middle tine, the high stress when interlocking on the distal tine suggests that the fighting was likely more constrained and predictable than among extant deer, likely involving twisting motions, as is known in extant deer with palmated antlers.[42]

In deer, gestation time generally increases with body size. A Шаблон:Cvt doe may have had a gestation period of about 274 days. Based on this and patterns seen in modern deer, last year's antlers in Irish elk bucks were potentially shed in early March, peak antler growth in early June, completion by mid-July, shedding velvet (a layer of blood vessels on the antlers in-use while growing them) by late July, and the height of rut falling on the second week of August. Geist, believing the Irish elk to have been a cursorial animal, concluded that a doe would have to have produced nutrient-rich milk so that her calf would have enough energy and stamina to keep up with the herd.[37]

Diet and life history

Файл:Irish Elk front.jpg
Skull in front view

The mesodont (meaning neither high (hypsodont) or low (brachydont) crowned) condition of the teeth suggests that the species was a mixed feeder, being able to both browse and graze. Pollen remains from teeth found in the North Sea around 43,000 years old were found to be dominated by Artemisia and other Asteraceae, with minor Plantago, Helianthemum, Plumbaginaceae and willow (Salix).[43] Another earlier specimen from the Netherlands (dating to the Eemian interglacial or early in the Last Glacial Period) was found to have pollen of Apiaceae, including cow parsley (Anthriscus sylvestris), cow parsnip/hogweed (Heracleum), water pennywort (Hydrocotyle), Asteraeceae, Filipendula, Symphytum and grass embedded with its teeth.[44] A stable isotope analysis of the terminal Pleistocene Irish population suggests a grass and forb based diet, supplemented by browsing during stressed periods.[45] Dental wear patterns of specimens from the late Middle and Late Pleistocene of Britain suggest a diet tending towards mixed feeding and grazing, but with a wide range including leaf browsing.[46]

Based on the dietary requirements of red deer, a Шаблон:Cvt lean Irish elk stag would have needed to consume Шаблон:Cvt of fresh forage daily. Assuming antler growth occurred over a span of 120 days, a stag would have required 1,372 g (3 lb) of protein daily, as well as access to nutrient- and mineral-dense forage starting about a month before antlers began sprouting and continuing until they had fully grown. Such forage is not very common, and stags perhaps sought after aquatic plants in lakes. After antler growing, stags could probably satisfy their nutritional requirements in productive sedge lands bordered by willow and birch forests.[37]

The Irish elk may have been preyed upon by the large carnivores of the time, including the cave lion,[47] and the cave hyena.[48]

Extinction

Outside of the Irish Late Pleistocene, remains of Irish elk are uncommon, suggesting that they were usually rare in the areas that they did occur.[4]

Historically, its extinction has been attributed to the encumbering size of the antlers, a "maladaptation" making fleeing through forests especially difficult for males while being chased by human hunters,[13] or being too taxing nutritionally when the vegetation makeup shifted.[33] In these scenarios, sexual selection by does for stags with large antlers would have contributed to decline.[49]

However, antler size decreased through the Late Pleistocene and into the Holocene, and so may not have been the primary cause of extinction.[40] A reduction in forest density in the Late Pleistocene and a lack of sufficient high-quality forage is associated with a decrease in body and antler size.[50] Such resource constriction may have cut female fertility rates in half.[40] Human hunting may have forced Irish elk into suboptimal feeding grounds.[3]

The distribution of M. giganteus is thought to have been strongly controlled by climactic conditions. The range of the Irish elk appears to have collapsed during the Last Glacial Maximum (LGM), with few remains known between 27,500 and 14,600 years ago, and none between 23,300 and 17,500 years ago. Known remains substantially increase during the latest Pleistocene, where it appears to have re-colonized most of its former range, with abundant remains in the UK, Ireland, and Germany.[4] A 2021 study found that M. giganteus saw a progressive decline in mitochondrial genome diversity beginning around 50,000 years ago, which accelerated during the LGM.[51]

While the range of the taxon was dramatically reduced after the Pleistocene-Holocene transition, it managed to survive into the middle Holocene in the central part of its range within European Russia and Western Siberia. It is suggested that extinction was contributed to by further climatic changes transforming preferred open habitat into uninhabitable dense forest.[4] The final demise may have been caused by several factors both on a continental and regional scale, including climate change and hunting.[50][52] The youngest dates in this region from Kamyshlov in Western Siberia and Maloarkhangelsk, Oryol Oblast In European Russia date to around 7,700-7,600 years ago, and it is suggested that it likely became extinct shortly after this time. Lister and Stewart concluded in a study of the extinction of the Irish elk that "it seems clear that environmental factors, cumulatively over thousands of years, reduced giant deer populations to a highly vulnerable state. In this situation, even relatively low-level hunting by small human populations could have contributed to its extinction."[4]

Relationship with early humans

Шаблон:Multiple image

A handful of Irish elk depictions are known from the art of the Upper Paleolithic in Europe. However, these are much less abundant than the common red deer and reindeer depictions. The bones of the Irish elk are uncommon in localities where they are found, and only a handful of examples of human interaction are known.[4] A mandible from Ofatinţi, Moldova dating to either the Eemian or the early Late Pleistocene, "is peculiar because it has ancient tool-made notches on its lateral side".[53] At the Riparo del Broion site in northern Italy, dating to 50-44,000 years ago, a tibia and metapodial bone display slicing marks as a result of butchery, likely by Neanderthals.[54] Several M. giganteus bones from the Chatelperronian levels of the Labeko Koba site in Spain are noted for bearing puncture marks, which have been interpreted as anthropogenic.[55] A terminal Pleistocene (13710-13215 cal BP) skull from Lüdersdorf, Germany is noted to have had the antler and facial part of the skull removed in a way unlikely to be due to natural causes.[56] A calcaneum from an associated lower hind limb from the early Holocene site of Sosnovy Tushamsky in Siberia is noted to have "two short and deep traces of cutting blows", which are interpreted as "clear evidence of butchery".[57][4] The use of shed antler bases is also known, at the terminal Pleistocene (Allerød) Endingen VI site in Germany, a shed antler base appears to have been used in a way analogous to a lithic core to produce "blanks" for the manufacture of barbed projectile tips.[58][4] A ring-like mark on a shed antler beam from the similarly aged Paderborn site in Germany has been suggested to be anthropogenic.[59]

Modern significance

Файл:Crystal Palace Megaloceros giganteus.jpg
Sculptures in Crystal Palace

Due to the abundance of Irish elk remains in Ireland, a thriving trade in their bones existed there during the 19th century to supply museums and collectors. Skeletons and skulls with attached antlers were also prized ornaments in aristocratic homes. The remains of Irish elk were of high value: "In 1865, full skeletons might fetch £30, while particularly good heads with antlers could cost £15." with £15 being more than 30 weeks' wages for a low skilled worker at the time.[60] Indeed Leeds Philosophical and Literary Society bought a full skeleton in 1847, from Glennon's in Dublin, for £38.[61] This specimen, discovered at Lough Gur near Limerick, is still on display at Leeds City Museum.[62]

A folk memory of the Irish elk was once thought to be preserved in the Middle High German word Шаблон:Lang, a large beast mentioned in the 13th-century Nibelungenlied along with the then-extant aurochs (Шаблон:Lang / "After this he straightway slew a Bison and an Elk, Of the strong Wild Oxen four, and a single fierce Schelch.") This opinion is no longer widely held.[13] The Middle Irish word Шаблон:Lang was also suggested as a reference to the Irish elk.[63][64] Turf cutters of Clooney and Tulla in County Clare, Ireland referred to the Irish elk as the Шаблон:Lang (Шаблон:Lang meaning Deer, Шаблон:Lang meaning "great", or "large").[65] However, these interpretations are not conclusive.[66]

See also

References

Шаблон:Reflist

Further reading

Kurten is a paleo-anthropologist, and in this novel, he presents a theory of Neanderthal extinction. Irish elk feature prominently, under the name shelk which Kurten coins (based on the aforementioned old German schelch) to avoid the problematic aspects of "Irish" and "elk" as discussed above. The book was first published in 1980 when "Giant Deer" was not yet being used widely.
  • Zoological Science 22: 1031–1044 (2005).
  • Larson, Edward J. (2004). Evolution: The Remarkable History of a Scientific Theory.

External links

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  1. ↑ Шаблон:Cite book
  2. ↑ 2,0 2,1 2,2 2,3 2,4 Шаблон:Cite journal
  3. ↑ 3,0 3,1 Шаблон:Cite journal Supplementary information. Erratum in Шаблон:Cite journal
  4. ↑ 4,0 4,1 4,2 4,3 4,4 4,5 4,6 4,7 4,8 Шаблон:Cite journal
  5. ↑ 5,0 5,1 5,2 5,3 5,4 5,5 Шаблон:Cite journal
  6. ↑ 6,0 6,1 6,2 6,3 Шаблон:Cite journal
  7. ↑ 7,0 7,1 Шаблон:Cite journal
  8. ↑ 8,0 8,1 8,2 Шаблон:Cite journal
  9. ↑ Шаблон:Cite journal
  10. ↑ 10,0 10,1 Blumenbach J. 1799. Handbuch der Naturgeschichte (6th Ed.) 16: 697
  11. ↑ 11,0 11,1 11,2 11,3 11,4 Шаблон:Cite journal
  12. ↑ Wright, T. 1748. Louthiana: Or, an Introduction to the Antiquities of Ireland; W. Faden: London, UK; p. 20
  13. ↑ 13,0 13,1 13,2 13,3 13,4 Шаблон:Cite journal
  14. ↑ Joshua Brookes (1827) "Brookesian Museum. The Museum of Joshua Brookes, Esq. Anatomical and Zoological Preparations" London Gold and Walton
  15. ↑ 15,0 15,1 15,2 15,3 Lister, A M, 1987 Megaloceros Brookes 1828 Mammalia Artiodactyla Proposed Emendation Of The Original Spelling The Bulletin of zoological nomenclature. 44 255–256
  16. ↑ Шаблон:Cite web
  17. ↑ Шаблон:Cite book
  18. ↑ International Commission on Zoological Nomenclature 1989. Opinion 1566. Megaloceros Brookes, 1828 (Mammalia, Artiodactyla): original spelling emended. Bulletin of zoological nomenclature 46: 219–220.
  19. ↑ Anderson, Kristina. "What On Earth - A Canadian Newsletter for the Earth Sciences". What On Earth - A Canadian Newsletter for the Earth Sciences. N.p., 15 November 2002. Web. 23 October 2014.
  20. ↑ 20,0 20,1 Zimmer, Carl. "The Allure of Big Antlers". The Loom. Discover, National Geographic. 3 September 2008. Web. 23 October 2014.
  21. ↑ Шаблон:Cite journal
  22. ↑ 22,0 22,1 22,2 22,3 22,4 22,5 22,6 Шаблон:Cite journal
  23. ↑ 23,0 23,1 23,2 23,3 23,4 23,5 Шаблон:Cite book
  24. ↑ Шаблон:Cite journal
  25. ↑ Шаблон:Cite journal
  26. ↑ 26,0 26,1 Шаблон:Cite journal
  27. ↑ Шаблон:Cite journal
  28. ↑ Шаблон:Cite web
  29. ↑ Шаблон:Cite journal
  30. ↑ 30,0 30,1 Шаблон:Cite journal
  31. ↑ Шаблон:Cite journal
  32. ↑ Van der Made, J. Giant deer. In Elefantenreich. Eine Fossilwelt in Europa; Meller, H., Ed.; Verlag Beier & Beran: Halle, Germany, 2010; pp. 408–412
  33. ↑ 33,0 33,1 33,2 33,3 33,4 33,5 Шаблон:Cite journal
  34. ↑ 34,0 34,1 R. D. E. Mc Phee, Extinctions in Near Time: Causes, Contexts, and Consequences p. 262
  35. ↑ A View to a Kill: Investigating Middle Palaeolithic Subsistence Using an Optimal Foraging Perspective
  36. ↑ Шаблон:Cite journal
  37. ↑ 37,0 37,1 37,2 37,3 37,4 37,5 37,6 Шаблон:Cite book
  38. ↑ Шаблон:Cite book
  39. ↑ 39,0 39,1 Шаблон:Cite journal
  40. ↑ 40,0 40,1 40,2 40,3 Шаблон:Cite journal
  41. ↑ Шаблон:Cite journal
  42. ↑ Шаблон:Cite journal
  43. ↑ Шаблон:Cite journal
  44. ↑ Шаблон:Cite journal
  45. ↑ Шаблон:Cite journal
  46. ↑ Шаблон:Cite journal
  47. ↑ Шаблон:Cite journal
  48. ↑ Шаблон:Cite journal
  49. ↑ Шаблон:Cite journal
  50. ↑ 50,0 50,1 Шаблон:Cite journal
  51. ↑ Шаблон:Cite journal
  52. ↑ "Irish Elk Survived after Ice Age Ended" Author(s): S. P. Science News, Vol. 166, No. 19 (6 November 2004), p. 301. Society for Science & the Public.
  53. ↑ Шаблон:Cite journal
  54. ↑ Шаблон:Cite journal
  55. ↑ A. Villaluenga, A. Arrizabalaga, J. Rios-Garaizar "Multidisciplinary approach to two Chatelperronian series: lower IX layer of Labeko Koba and X level of Ekain (Basque Country, Spain)" Journal of Taphonomy, 10 (2012), pp. 499–520
  56. ↑ B. Bratlund "Ein Riesenhirschschädel mit Bearbeitungsspuren aus Lüdersdorf", Kreis Grevesmühlen. Offa 49/50 (1992/1993) (1994), pp. 7–14
  57. ↑ S. K. Vasiliev, V. S. Slavinsky, A. V. Postnov. "The Irish elk (Megaloceros giganteus Blumenbach, 1803) in the paleofauna of the Holocene sites of the northern Angara region (Ust-Tushama-1, Sosnovy Tushamsky Ostrov, Ust-Talaya)" Journal of Novosibirsk State University, Series: Hist. Philol., 12/7 (2013), pp. 177–185 Archaeology and Ethography
  58. ↑ K. Kaiser, P. De Klerk, T. Terberger. "Die 'Riesenhirsch Fundstelle' von Endingen: geowissenschaftliche und archäologische Untersuchungen an einem spätglazialen Fundplatz in Vorpommern". Eiszeitalt und Gegenwart. 49 (1999). pp. 102–123.
  59. ↑ M. Baales, S. Birker, H.-O. Pollman, W. Rosendahl, B. Stapel "Erstmals datierte organische Artefakte aus dem Spätpaläolithikum Westfalens Archäologie in Westfalen". Lippe, 2012 (2012), pp. 24–27
  60. ↑ Шаблон:Cite journal
  61. ↑ Letter from R. Glennon, Dublin, to Henry Denny, Leeds, 10 July 1847. Held at Leeds Discovery Centre, Leeds, UK.
  62. ↑ Шаблон:Cite web
  63. ↑ Шаблон:Cite book
  64. ↑ Шаблон:Cite journal
  65. ↑ Шаблон:Cite web
  66. ↑ Шаблон:Cite book
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